REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN
MALESIA
ABDULROKHMAN KARTONEGORO
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
LETTER OF STATEMENT
I express that thesis entitled:
REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN MALESIA is true represent result of my own research and have never been published. All information and data that used have been expressed clearly and can be checked its truth.
Bogor, June 2011
ABSTRACT
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
Revision on Rhynchoglossum in Malesia has been done. This research conducted based on morphological characters of 233 sheets of Herbarium specimen and Herbarium specimen photographs from Herbarium Bogoriense (BO), Singapore Botanic Gardens (SING), National Herbarium Netherlands (L), Royal Botanic Garden Kew (K), Natural History Museum London (BM), Royal Botanic Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian Institute (US), and Harvard University (A). The result shows that six species are recognized in Malesia and five are endemic. Five species are known previously namely R. borneense, R. klugioides, R. medusothrix, R. obliquum, and
R. spumosum; one species is proposed as a new and newly described namely
R. celebicum. Rhynchoglossum borneense and R. medusothrix endemic to Borneo,
R. celebicum endemic to Sulawesi, while R. spumosum endemic to Philippine Islands. Rhynchoglossum klugioides and R. obliquum distributed wider in Malesia. Rhynchoglossum papuae and R. merrilliae are placed under synonym of
R. obliquum and R. spumosum. A phylogenetic analysis using PAUP version 4.0 program based on 60 morphological characters with Stauranthera coerulea and
Loxonia hirsuta as an outgroup was undertaken. The analysis resulted a parsimonious tree which shown that Rhynchoglossum in Malesia can be divided in two groups. First clade consist R. obliquum, R. spumosum and R. celebicum while the second clade consist R. klugioides, R. borneense and R. medusothrix. Taxonomic treatment was giving with a identification key to the species, description, nomenclature, distribution, and notes each species.
Keywords: Gesneriaceae, Malesia, phylogenetic analysis, revision,
ABSTRAK
Abdulrokhman Kartonegoro. Revisi Rhynchoglossum (Gesneriaceae) di Malesia. Dibimbing oleh Dr. Sri S. Tjitrosoedirdjo, M.Sc. dan Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
Revisi marga Rhynchoglossum di Malesia telah dilakukan. Studi ini dilakukan berdasarkan pendekatan karakter morfologi pada 233 lembar spesimen herbarium dan foto spesimen herbarium dari Herbarium Bogoriense (BO), Singapore Botanic Gardens (SING), National Herbarium Netherlands (L), Royal Botanic Garden Kew (K), Natural History Museum London (BM), Royal Botanic Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian Institute (US), dan Harvard University (A). Hasil revisi menunjukkan ada enam jenis yang diketahui di Malesia dan lima jenis endemik. Lima jenis merupakan jenis-jenis yang sudah dikenal sebelumnya yaitu R. borneense, R. klugioides, R. medusothrix,
R. obliquum, dan R. spumosum; satu jenis diusulkan sebagai jenis baru dan baru dipertelakan yaitu R. celebicum. Rhynchoglossum borneense dan R. medusothrix
endemik di Borneo, R. celebicum endemik di Sulawesi, sedangkan R. spumosum
endemik di Filipina. Rhynchoglossum klugioides dan R. obliquum tersebar lebih luas di Malesia. Rhynchoglossum papuae dan R. merrilliae dimasukkan ke dalam sinonim dari R. obliquum dan R. spumosum. Analisis filogenetik menggunakan program PAUP versi 4.0 berdasarkan pada 60 karakter morfologi dengan
Stauranthera coerulea dan Loxonia hirsuta sebagai outgroup. Dari hasil analisis dihasilkan pohon parsimoni yang menunjukkan bahwa Rhynchoglossum di Malesia dapat dipisahkan menjadi dua kelompok. Kelompok pertama terdiri atas
R. obliquum, R. spumosum dan R. celebicum sedangkan kelompok kedua terdiri atas R. klugioides, R. borneense dan R. medusothrix. Bagian taksonomi disampaikan dengan kunci identifikasi jenis, pertelaan, tata nama, distribusi dan catatan tiap-tiap jenis.
SUMMARY
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
Rhynchoglossum Blume is a genus of fleshy herb belongs to family
Gesneriaceae. It is comprised about 10 species all over the world. The genus
Rhynchoglossum was established by Blume with one spesies of Rynchoglossum obliquum. Rhynchoglossum can be recognized morphologically as fleshy herb plant with anisophyllous decussate leaf or alternate leaf arrangement, asymmetrically leaf blade and the unilaterally inflorescence. The genus has a little economic value but until now still planted almost 130 years in several Botanic Gardens and private gardens around the world.
Several taxonomic study of Rhynchoglossum has done since the genus proposed by Blume. From many studies and revisions, there was no specific taxonomic revision for Malesian region that known as the center of biodiversity of the genus. Revision on this genus in Malesia has not been done before. Based on that information, the revision of the genus Rhynchoglossum is needed.
The research objective of this study was to revise taxonomic of
Rhynchoglossum in Malesia. Taxonomic revision was done to find the information about generic and species delimitation, by providing data on its diversity, identification key for species, analytic description for genus and species, distribution pattern, and phylogenetic of Rhynchoglossum in Malesia.
University (A), Smithsonian Institute (US), and New York Botanical Gardens (NY) were also studied. Phylogenetic analysis of Malesian Rhynchoglossum was studied based on sixty morphological characters which collected during morphological observation and analyzed based on Maximum Parsimony using PAUP version 4.0b.10 with the Heuristic search settings vegetative and floral characters were selected for the analysis.
Based on the revision of Rhynchoglossum in Malesia, there were six species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B.L.Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no infraspecific status decided in this revision. Rhynchoglossum celebicum is new species that newly described here beside the other five species.
In Malesia this genus consist of six species and five of them are endemic to the region. Rhynchoglossum klugioides was previously only known from Philippines now distributed also to Seram Island, Moluccas. Four species are known endemic in each region. Rhynchoglossum spumosum is endemic to Philippines known from Negros and Mindanao, while the R. celebicum is endemic to Sulawesi. Rhynchoglossum borneense and R. medusothrix known endemic to Borneo which is known only found in the eastern part of the island.
Rhynchoglossum can be found from lowland areas to mountain areas of 1500 m above sea level at the edge of the forest, close to water shaded or in the rock of limestone area.
The phylogenetic analysis using 60 morphological characters based on Maximum Parsimony method. The 128 equally most parsimonious tree of 138 steps produce consistency index (CI) = 0.6562, homoplasy index (HI) = 0.3438 and retention index (RI) = 0.5769. The higher CI value was shown that the number of homoplasy rather low. The separation of the ingroup and outgroup was supported with a high bootstrap value. The ingroup was separated in two sister clades, one consist R. obliquum, R. spumosum and R. celebicum, and the other consist of R. klugioides, R. borneense and R. medusothrix.
Keywords: Gesneriaceae, Malesia, phylogenetic analysis, revision,
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REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN
MALESIA
ABDULROKHMAN KARTONEGORO
Thesis submitted
As partial fulfillment requirement for the Master Degree In Plant Taxonomy
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
Title : Revision of Rhynchoglossum (Gesneriaceae) in Malesia Name : Abdulrokhman Kartonegoro
NRP : G353080061 Study Program: Plant Biology
Approved by Supervisor Committee
Dr. Sri S. Tjitrosoedirdjo, M.Sc. Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
(Chairman) (Member)
Head of Plant Biology Study Program Dean of Graduate School
Dr. Ir. Miftahudin, M.Si. Dr. Ir. Dahrul Syah, M.Sc. Agr.
ACKNOWLEDGEMENT
I would like to thank to my supervisors, Dr. Sri S. Tjitrosoedirdjo (IPB) and Prof.(R) Dr. Elizabeth A. Widjaja (LIPI) for their valuable advices and guidance in completing this thesis. I am gratefully thanked to Dr. Teguh Triono (BO), Prof. Dr. Eko Baroto Walujo (BO), Dr. Harry Wiriadinata (BO), W.S. Hoover (NETC) for their helpful giving advises for completing this thesis. I also would like thank to Dr. David Middleton (E), Dr. Laurence Skog (US), Dr. Ruth Kiew (KEP), and Dr. John R. Clark (SEL) for their helpful advices and critics to my thesis draft especially in Gesneriaceae. Further thanked to Dr. J.F. Veldkamp for helping in nomenclature taxonomy and latin diagnose for the new species.
Special thank to The Indonesian Botanical Exploration and Taxonomy Project (IBETP) of New England Tropical Conservatory (NETC), Nellie Sleeth Scholarships Grant of Gesneriad Society USA and Singapore Botanic Gardens Fellowship Grant for financing my research and study at the Graduate School of Bogor Agricultural University (IPB).
I would like to express my gratitude to Herbarium Bogoriense (BO) of Research Center for Biology (LIPI) and Singapore Botanic Gardens for granting permission to conduct my research work and providing some facilities. I would like to thanks especially to the Director of Herbaria A, BO, BM, E, L, K, SING, US, and NY for examined their valuable specimens. Special thanks giving to my friends Lim Chung Lu and Yao Tze Liong from Forest Research Institue Malaysia and Mr. Gerard Thijsse from National Herbarium Netherland for sending some image of Rhynchoglossum Type from Kew and Leiden.
I thanks to my colleagues and friends in Herbarium Bogoriense and Plant Biology Major IPB especially Alex Sumadijaya, Ina Erlinawati, Yessi Santika, Arief Hidayat, Destri, Wina Dyah Puspitasari, Sukiman, Rina, Adeel, Sanou Faye for their help, support and opportunity to cement a good friendship.
CURRICULUM VITAE
The author was born in Jakarta on 4 January 1980, the fifth son of five of Mr. Bunyamin (Alm) and Mrs. Muhanah.
In 1999, he became a student at the Department of Biology, Faculty of Mathematic and Natural Science, University of Indonesia, Depok and graduated at 2005.
Since April 2006 he was employed by the Research Center for Biology, Indonesian Institute of Sciences (LIPI) in Cibinong, West Java and still active till now.
LIST OF TABLES
LIST OF FIGURES
6 Cladogram of phylogenetic tree of Rhynchoglossum in Malesia.Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769.
The number shown above line showed supported the Bootstrap values …. 22 7 Rhynchoglossum borneense Merr., Elmer 21467 from Tawao, Malaysian Borneo (isotype SING) ...………...………... 29 8 Distribution map of Rhynchoglossum borneense in Malesia in the circle …. 30 9 Distribution map of Rhynchoglossum celebicum in Malesia in the circle ... 31 10 Rhynchoglossum celebicum Karton., Eyma 1572 from Maraowa, Sulawesi (holotype BO) ………...……… 32 11 Distribution map of Rhynchoglossum klugioides in Malesia in the circle .... 34 12 Rhynchoglossum klugioides C.B.Clarke, Cuming 824 from Tayabas, Luzon (holotype K) ……….. 35 13 Distribution map of Rhynchoglossum medusothrix in Malesia in the circle .. 37 14 Rhynchoglossum medusothrix B.L.Burtt, Kostermans 13994 from Berau, East Kalimantan, Borneo (isotype BO) .……… 38 15 Distribution map of Rhynchoglossum obliquum in Malesia in the circle ... 43 16 Rhynchoglossum obliquum Blume, Blume 52 from Java (lectotype L) …. 44 17 Distribution map of Rhynchoglossum spumosum in Malesia in the circle .. 46 18 Rhynchoglossum spumosum Elmer, Elmer 9929 from Cuernos Mts.,
LIST OF APPENDICES
Page 1 List of species Rhynchoglossum proposed in Malesia (Burtt 1962) …... 57 2 Matrix of 60 Morphological Characters for Maximum Parsimony
INTRODUCTION
Rhynchoglossum Blume was a genus of fleshy herb belongs to family
Gesneriaceae. It comprised of 10 species all over the world. It is classified in the tribe Epithemateae or EpithematoidGesneriaceae together with Epithema Blume,
Gyrogyne W.T. Wang, Loxonia Jack, Monophyllaea R. Br., Stauranthera Benth., dan Whytockia W.W. Sm. Rhynchoglossum was mainly found from India, Ceylon, Nepal, South China, Formosa, Indochina, Thailand, and Malesia, with only one species grow in Southern America (Mayer et.al. 2003; Weber 2004b).
The genus Rhynchoglossum was established by Blume (1826) with one spesies of Rynchoglossum obliquum. The type specimen was collected from Java. The species has a wide distribution from India, Ceylon, South China, Indochina, and Malay Archipelago through New Guinea. Some genus allied like Antonia R. Br., Glossanthus Benth., Klugia Schltdl., and Loxotis R. Br. are proposed later as the synonym of Rhynchoglossum (Burtt 1962).
Rhynchoglossum can be recognized morphologically as fleshy herb plant with anisophyllous decussate leaf or alternate leaf arrangement, asymmetrically leaf blade and the unilaterally inflorescence. The genus has a little economic value but several Botanic Gardens and private gardens are growing the plant as an ornamental (Skog 1985). One of the most ornamental planted is Rhynchoglossum gardneri Theobald & Grupe from India and Ceylon. The epithet name of
Rhynchoglossum is from Greek whereas Rhynchos means beak; and glossa means tongue. The second part of the name clearly alludes to the broad, tongue-like lower lip of the corolla, the first part perhaps to the narrow corolla tube or to the pointed petal tips (Weber 2004b).
Weber (2004b) put Rhynchoglossum under Epithematoid Gesneriaceae. This group has unique and special characters of unequal cotyledons, asymmetrically leaf blade and capsule fruit. The distribution of the group was mostly in tropical Asia, few from Africa, and one species from southern America. The disjunction distribution from this group was very unique especially in
Several taxonomic study of Rhynchoglossum has done since the genus proposed by Blume. Most of the studies generally observed the family
Gesneriaceae, whereas the Rhynchoglossum was included. Some of comprehensive studied were by Bentham (1876) and Clarke (1883). The latest study was a little revision done by Burtt (1962) which is stated that
Rhynchoglossum has 13 species and with very limited material specimens. From many studies and revisions, there was no specific taxonomic revision for Malesian region that known as the center of biodiversity for the genus. Revision about species biodiversity and the number of species in Malesia has not been done before. Based on that information, the revision of the genus Rhynchoglossum is needed.
The research objective of this study was to revise taxonomic of
LITERATURE REVIEW
Rhynchoglossum taxonomically included in Epithematoid Gesneriaceae
group or tribe Epithemateae along with others genera like Whytockia W.W. Sm.,
Gyrogyne W.T. Wang, Epithema Blume, Monophyllaea R. Br., Loxonia Jack, and
Stauranthera Benth. (Weber 2004b). Generally these groups have a similarity in unequal cotyledons, unequal leaf arrangement in one node, asymmetrically leaf blade and capsule fruits. Rhynchoglossum is allied with Loxonia and Stauranthera
in the shape of leaf arrangement. Rhynchoglossum has similarity with
Monophyllaea in the shape of inflorescences with unilateral cymose shape (Weber 2004b).
In the earlier status, Rhynchoglossum distributed from India, Ceylon, China, Taiwan, Indochina to New Guinea in the tropical Asia (Weber 2004b). Burtt (1962) included Klugia Schltdl. as synonymy of Rhynchoglossum, which is
Klugia has a distribution in Southern America from Mexico to Peru. The united of
Klugia as a synonym makes distribution of Rhynchoglossum wider and disjunct to America (Figure 1). These disjunction distributions on the genus within Australasia to America make Rhynchoglossum very unique and have an interesting distribution history. Burtt (1998) give preliminary hypothesis that
Rhynchoglossum reach America from Asia via Africa from where it now has completely disappeared or no species are found.
Figure 1 Distribution of Rhynchoglossum in yellow circle
Rhynchoglossum obliquum Blume is the common and widespread species, found from India, China to Malesia (Bakhuizen van den Brink Jr. 1965; Burtt 1962). In the Southern America only have one species R. azureum (Schltdl.) B.L. Burtt which has disjunctions distribution with its allies in Asia especially in Southern India (Mayer et.al. 2003). Some of the other species (e.g. R. borneense
Merr., R. medusothrix B.L.Burtt, and R. spumosum Elmer) only have restricted distributions and even with endemic pattern. Its preferably habitat is on the wet and shady (especially limestone) rocks, in the forest or open vegetation or shady places; usually grow in the lowlands (Weber 2004b).
strongly zygomorphic; tube cylindrical, white; limb strongly bilabiate, upper lip of two short, upright or reclined lobes, lower lip a large, roundish or elongate, not or only slightly three-partite tongue of azure colour. Fertile stamens four (the former genus Klugia) or two (Rhynchoglossum s.str.). Ovary globose-ovoid. Fruit a globose or ovoid capsule included in the calyx, dehiscing loculicidally by 2 valves (Weber 2004b). The illustration of the genus show on Figure 2.
Figure 2 Illustration of Loxotis obliqua = Rhynchoglossum obliquum
Brown (1832) proposed a new genus of Antonia with the species of
A. obliqua (Wall.) R. Br. which is combination of Wulfenia obliqua Wall. (Weber 2004a). Three years later Brown moved the species into the genus Loxotis R. Br. ex Benth. and made a new combination of L. obliqua (Wall.) R. Br.. The species is surviving for a long time as well as Miquel (1855) use it. Later on Clarke (1883) realizes that Antonia and Loxotis are superfluous because it is similar and identical to Rhynchoglossum and then he put both genera under synonym of
Rhynchoglossum. Recently Burtt (1962) also included Klugia to a synonym because it is only a variation of Rhynchoglossum.
The latest study by Burtt (1962) sign 13 species of Rhynchoglossum all over the world comprising 10 species from Asiatic and 3 from America. After that the number species of Rhynchoglossum are decrease because reducing some species into synonym. Rhynchoglossum grandiflorum (Fritsch) B.L. Burtt and
R. violaceum (Fritsch) B.L. Burtt are turn into synonym of R. azureum because of the similarity in morphology and geography (Wiehler 1983). Wang et.al. (1998) reduce R. hologlosum Hayata into synonym of R. obliquum. Since the reduction of some species into synonim, total number species of Rhynchoglossum known only 10 species all over the world (Mayer et.al. 2003; Weber 2004b).
History of the number species proposed in Malesia start when Blume (1826) established the R. obliquum as the type of the genus from Java. After Blume published the type then followed by de Candolle (1845) which established another species R. blumei DC. from Java. Clarke (1883) included R. blumei to be a synonym of R. obliquum and established a new species of R. klugioides C.B. Clarke from Luzon, Philippines. The species epithet is given by Clarke as a species which have similarity with Klugia. In the early of twentieth century, Elmer (1908) proposed a new R. spumosum Elmer from Negros Philippines and followed by Kraenzlin (1913) proposed R. merrilliae Kraenzl. from Mindanao, Philippines. Ten years later, a new species of Rhynchoglossum proposed by Schlechter (1923) which is from New Guinea and named as R. papuae Schltr. Another species from Borneo proposed by Merrill (1929) and named as
proposed by Burtt (1962) also from Borneo and completely sign seven species number in Malesia (Appendix 1).
Phylogenetic analysis of Rhynchoglossum using molecular DNA was studied together with another member of tribe Epithemateae. Phylogenetic tree showed that Rhynchoglossum separate in different clade with another
Epithemateae and act as sister position. This separation is in accordance with many and strong morphological differences (e.g., alternate leaf arrangement, strongly asymmetrical leaves, terminal inflorescences in the form of unilateral racemes, enlarged lower lip of corolla). The genus has many special characters in common with the other Epithemateae, except Monophyllaea and Whytockia. In the Rhynchoglossum group reflected in the phylogram by two clades;
MATERIALS AND METHODS
This study was carried out at the Systematic Laboratory of the Herbarium Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI) at Cibinong, West Java and the Herbarium of Singapore Botanic Gardens, Singapore National Parks from January 2010 to February 2011. It is based on 233 sheets of Herbarium specimens (including spirit collections) of Rhynchoglossum
from the Herbarium Bogoriense (BO) and the Herbarium of Singapore Botanic Gardens (SING). A number of herbarium specimen photographs from National Herbarium of Nederland-Universiteit Leiden Branch (L), Natural History Museum of London (BM), Royal Botanic Garden of Kew (K), Royal Botanic Garden of Edinburgh (E), Harvard University (A), Smithsonian Institute (US), and New York Botanical Gardens (NY) were also studied.
The materials were studied based on morphological characters using 10 x 40 binocular microscope and stereo microscope. The information was taken under the information data of location, general habitat, habit, stem, leaves, inflorescences and infructescences. Morphological description was following Veldkamp (1987). The revision methods were based on Rifai (1976, 2008) and Vogel (1987).
Table 1 The characters and characters state used in the phylogenetic analysis 6 Leaf shape ovate-elliptic (0); oblong (1)
7 Lamina membranous (0); subcoriaceous (1) 8 Leaf color pale green (0); dark green (1)
9 Leaf apex acuminate (0); acute(1) 10 Apex tip < 1 cm (0); 1 cm (1)
11 Leaf margin entire (0); dentate or serrate (1)
12 Leaf base rounded-cuneate (0); rounded-angustate (1); cordate-cuneate (2); cordate-angustate (3) 13 Midrib indument glabrous (0); puberulous (1); pubescent (2) 14 Abaxial leaf indument glabrous (0); glabrescent (1); pubescent (2) 15 Petiole indument glabrous (0); glabrescent (1); pubescent (2) 16 Petiole length < 10 cm (0); 10-15 cm (1); > 15 cm (2) 17 Inflorescence position axillar only (0); axillar and terminal (1) 18 Inflorescence growth erect (0); spreading (1)
19 Terminal infl. length ≤ 15 cm (0); > 15 cm (1) 20 Axillar infl. length ≤ 5 cm (0); > 5 cm (1)
21 Inflorescence side multilateral (0); unilateral (1) 22 Flower number ≤ 10 (0); > 10 (1)
23 Main axis indument glabrous (0); puberulous (1); pubescent (2) 24 Peduncle length ≤ 3 cm (0); > 3 cm (1)
25 Bract shape linier (0); oblong (1) 26 Bract length ≤ 4 mm (0); > 4 mm (1)
27 Pedicel indument glabrous (0); glabrescent (1); puberulous (2); pubescent (3)
Table 1 continued
NO CHARACTERS CHARACTER STATE
28 Pedicel length ≤ 3 mm (0); > 3 mm (1) 29 Bracteole length ≤ 5 mm (0); > 5 mm (1)
30 Bracteole position rise from base pedicel (0); rise from middle pedicel (1)
31 Flower length ≤ 15 mm (0); > 15 mm (1) 32 Calyx tube shape campanulate (0); urceolate (1)
33 Calyx tube color white (0); pale green (1); dark green (2) 41 Adaxial lip shape 2-lobed (0); emarginated (1) 42 Adaxial lip length ≤ 3 mm (0); > 3 mm (1) 43 Abaxial lip shape tongue-like (0); 3-lobed (1) 44 Abaxial lip apex rounded (0); obtuse (1) 45 Abaxial lip length ≤ 5 mm (0); > 5 mm (1) 46 Stamens number 2 (0); 4 (1)
47 Smaller stamen fil. shape terete (0); flat (1) 48 Smaller stamen fil. length ≤ 2 mm (0); > 2 mm (1)
Table 1 continued
NO CHARACTERS CHARACTER STATE
55 Style length ≤ 10 mm (0); > 10 mm (1)
56 Fruit shape globose (0); ovoid (1); ellipsoid (2); capsule-like (3); elongate (4)
57 Stalk length ≤ 10 mm (0); > 10 mm (1)
58 Stalk indument glabrous (0); puberulous (1); pubescent (2) 59 Seed surface tessellate (0); smooth (1)
60 Seed length ≤ 0.4 mm (0); > 0.4 mm (1)
RESULT AND DISCUSSION
Based on this revision work of Rhynchoglossum in Malesia, there were six species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton. spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B. L. Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no infraspecific status decided in this revision. Rhynchoglossum celebicum is one of new species that newly described here beside the other five species.
New status has been given for some species in this revision.
Rhynchoglossum papuae Schltr. from New Guinea now turn into synonym of
R. obliquum. Another species is R. merrilliae Kraenzl. from Philippine now under synonym of R. spumosum. This new status decreasing the species number of
Rhynchoglossum in Malesia that giving by Burtt (1962) from seven to six here with new species that newly described (R. celebicum).
Based on preliminary revision, Burtt (1962) proposed a seven species of
Rhynchoglossum comprising R. borneense, R. klugioides, R. medusothrix,
R. merrilliae, R, obliquum, R. papuae, and R. spumosum (Appendix 1). In this recent revision, the number species are decreased because the status of two species changed. Rhynchoglossum merrilliae is synonym of R. spumosum due to its similarity on their morphological characters such as dwarf habit, small leaf blade, few flower in the inflorescence and two fertile stamens. Based on the observation to type specimens, it is realized that both are actually one species and
R. spumosum is used because this species name used earlier.
Rhynchoglossum papuae was proposed by Schlechter (1923) based on the small corolla lip and globose or ovoid capsules comparing with R obliquum. When more specimens were examined from New Guinea in this revision, it is seen that most characters from each specimen have no different with R. obliquum
new species (R. celebicum) in this revision, it showed that the total number of
Rhynchoglossum in Malesia become six species.
General Morphology of Rhynchoglossum in Malesia
Habit
Rhynchoglossum are erect and creeeping fleshy-succulent herbs plant from 5-150 cm height. The roots are adventitious with rhizomatous or non-rhizomatous. The only rhizomatous species is R. spumosum is being creeping in habit and dwarfing about 5-8 cm height. The stems usually are terete and wrinkled when dried. They have a variable indument, the hairs being absent or simple puberulous to pubescent. Most of the species have glabrous and smooth stem surface and have swollen nodes.
Rhynchoglossum in broad sense have a growing habit as an herb with perennial or annual growth. All Malesian species known have annual growth with fleshy succulent herb. Rhynchoglossum spumosum is the only species have creeping habit with rhizomatous root system. This habit makes the species has only maximum height 8 cm known as a dwarf Rhynchoglossum. Another species that reporting have rhizomatous habit is R. omeiense W.T.Wang from China, but different in having perennial growth (Wang et.al. 1998). Because of that it is necessary to study further by their molecular systematic to understand the uncontinous charcters occurs in several species.
Leaves
Leaf arrangements are alternate and exstipulate. The leaf blade of all
blades texture are mostly membranous, only R. klugioides have subcoriaceous leaf blade. Most of the species have glabrous and smooth leaf surface. Some have pubescent or puberulous indumentum when juvenile.
A B
C
Figure 3 Leave shape. A. Elliptic of R. obliquum B. Ovate of R. klugioides C. Oblong of R. celebicum
Rhynchoglossum have alternate leaf arrangement with no stipules. Different from other Gesneriaceae which have opposite leaf arrangement Rhynchoglossum
in leaf arrangement with anisophylly leaf arrangement, not reducing like
Rhynchoglossum. The opposite sides in the leaf arrangement are usually free or with the inflorescences.
Most of the species of Rhynchoglossum known have ovate to elliptic leaf shape. In this revision a new character of leaf which is oblong belongs to
R. celebicum. This different on its leaf characters made R. celebicum distinct and realized as a new species. The leaf texture of Rhynchoglossum has two types, one usually membranous (most of Malesian species) and the other one is subcoriceous (R. klugioides). The subcoriaceous texture is related with the species that formerly belongs to Klugia (known as R. azureum and R. notonianum recently).
Inflorescences
The inflorescence in Rhynchoglossum is raceme with terminally and axillary grow. The axillary inflorescence are usually opposite with leaf. The inflorescences are spreading or secundiflorous which is not erect (Figure 4A). Number of flower in inflorescence varies from less than 3 in R. spumosum to 50 in R. obliquum. Flowers in inflorescence are unilateral because the reduction of other side flowers and arrange in two rows. The bracts usually absent in R. spumosum or occur in linier shape in R. obliquum. Sometimes in inflorescences have bracts that not supported flower or known as sterile bracts. Bracteoles in Rhynchoglossum are mostly linier which rise from base of pedicel in R. klugioides and R. spumosum; and rise from the middle of pedicel in other species.
A B
Figure 4 Rhynchoglossum obliquum A. Inflorescence; B. Flowers
Flowers
The flowers are zygomorphic, tube-like and vary from less than 1 cm long till more than 3 cm long. Calyx tubes are infundibuliformis with campanulate shape in most species or urceolate shape in R. klugioides. The calyx adnate at base sometimes winged with line fushion; white to green colour. Calyx lobes are triangular and entire, glabrous, 5-merous. Corolla tubes are glabrous to puberulent, with tube and limb. The limb has two lipped which is from the petal fused (Figure 4B). The adaxial lip usually smaller with two lobes and the abaxial lip larger 3-lobed or undivided bearing a rounded elongate or tongue–like. The petals usually have a bluish to dark purple color and a few is white. Some species like R. borneense, R. medusothrix and R. klugioides have a yellow dot pilose in the throat of abaxial limb. The pilose dot of limb in R. medusothrix usually being medussoid and dense.
Two enlarged lips corolla with enlarge lower one in the flower of
Rhynchoglossum are common in the genus. The variation among the species is usually shown in the lower lip characters. The species that having small flower or corolla (R. obliquum, R. spumosum and R. celebicum) have tongue-like lower lips and the species with large flower or corolla (R. klugioides, R. borneense and
Rhynchoglossum in Malesia have 2 or 4 stamens. Rhynchoglossum obliquum, R. klugioides, R. celebicum and R. spumosum have 2 stamens. Otherwise, R borneense and R. medusothrix have 4 stamens. All stamens in flowers are arranged coherent in pairs. The stamens are adnate to corolla tube in adaxial limb if only 2 stamens or in abaxial limb if there 4 stamens. The anthers are basifixed, thecae parallel and dehiscing longitudinally. Filaments are usually flat. The ovaries in most Rhynchoglossum are ovoid with one loculed. Stigma are capitates or undivided and the style are usually glabrous and still remnant in fruiting time.
Four numbers of stamens in Malesian Rhynchoglossum allied with another species of Rhynchoglossum (R. notonianum (Wall.) B.L.Burtt and R. azureum
(Schltd.) B.L.Burtt) was previously from the genus Klugia. The different with the species outside Malesia is in the arrangement of the stamens. R. notonianum and
R. azureum have four equal stamens that coherent together while R. borneense
and R. medusothrix have four unequal stamens (two larger and other two smaller stamens). The arrangement of the stamens is also coherent but in pair not together (dydinamous).
Fruits
The fruits in all Rhynchoglossum are capsule type with one loculed. The shape of the fruits may vary from capsule-like, ovoid to elongate with prominent calyx remnants (Figure 5). The calyx remnants are half closing the fruit in
R. celebicum (Figure 5C) and fully closing the fruit in other species (Figure 5A & B). The fruits are dehiscing loculicidally to base with two valvate, not twisted. The seeds are cuneate and unappendaged.
The shape of the fruits of Rhynchoglossum is varying. The ovoid fruit shape is known oly from R. obliquum, R. borneense, R. medusothrix and R. spumosum with calyx remnant closing fully the fruit and persistent.
A B
C
Figure 5 Fruit shape A. Ovoid of R. obliquum B. Elongate of R. klugioides
C. Capsule-like of R. celebicum
Distribution
Rhynchoglossum approximately have 10 species of fleshy herbs that found throughout South Asia, China, South East Asia and Malesia to Southern America (Weber, 2004). The Malesian region has about six species and five of them are endemic to the region (Table 2). Rhynchoglossum obliquum is a widespread and very common species distributed from Sumatra to New Guinea. Rhynchoglossum klugioides was previously only known from Philippines currently distributed also to Seram Island, Moluccas. The other four species are endemic in each region.
borneense and R. medusothrix are endemic to Borneo which is recorded only found in the eastern part of the island.
Table 2 Distribution of Rhynchoglossum in Malesia
Species Sumatra Peninsular
Based on the distribution pattern, the center of the diversity
Rhynchoglossum in Malesia is found in Borneo, Philippine and Sulawesi. Borneo has three species (R. borneense, R. medusothrix, and R. obliquum), Philippine has three species (R. klugioides, R. spumosum, and R. obliquum) while Sulawesi has two species (R celebicum and R. obliquum). The regions shows the high number of Rhynchoglossum species found comparing to the other islands like Sumatra, Peninsular Malaya, Java, Lesser Sunda, Moluccas and New Guinea. The endemicity of Rhynchoglossum in the center of diversity is also high. Beside
R. obliquum, the rest five species known endemic in that area while a few
R. klugioides also observed in Moluccas.
The phytogeography pattern for Rhynchoglossum is still related with the pattern of all Asiatic Rhynchoglossum. Burtt (1998) said that Gesneriaceae is a family of Gondwanic origin. Epithematoid Gesneriaceae which is
Rhynchoglossum inside is a relict group that was once much larger and had much wider distribution. The ancient of Rhynchoglossum is known from Asiatic mainlad which now has the most species occur. Based on that hypothesis is predicted that the Malesian species are spreading from west Wallace line to east line. The species that found in the east Malesia are R. obliquum and R. klugioides
which known from Moluccas and New Guinea. In fact, both species also known distributed in the west of Malesia like Sumatra, Java, Borneo and Philippine.
Rhynchoglossum in Malesia made the genus is typically Sunda shelf or West Malesia taxon.
Rhynchoglossum can be found from lowland areas to mountain areas of 1500 m above sea level at the edge of the forest, close to water shaded or in the rock of limestone area. Most of the species like R. obliquum, R. celebicum,
R. klugioides and R. spumosum found in the forest with canopy or edge forest with wet condition near the river. Rhynchoglossum borneense and R. medusothrix are known from the limestone habitat in Borneo. It is looks that the preferable habitat impacts the distribution of species.
Phylogenetic analysis
The phylogenetic relationships of the genus Rhynchoglossum in Malesia was analyzed in the present study based on morphological characters. Sixty characters were used in this analysis (Table 1). The sixty characters were selected for their apparent taxonomic values that are good to separate one taxon with the others.
Figure 6 Cladogram of phylogenetic tree of Rhynchoglossum in Malesia. Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769. The number shown above line showed supported the Bootstrap values.
The separation of the ingroup and outgroup in the phylogenetic analysis was supported with a high bootstrap value, 100 % based on some characters: growing habit (character 1), habit (character 2), leaf arrangement (character 5), leaf margin (character 11), inflorescence growth (character 18), inflorescence side (character 21), corolla tube color (character 37), fruit shape (character 56), and seed surface (character 59). The outgroups (Stauranthera coerulea (Blume) Merr. and Loxonia hirsuta Jack) are the closest taxa to the genus together in the tribe
Epithemateae. This group is monophyletic with Rhynchoglossum in the sister group with the other member of the tribe is in accordance with the many and strong morphological different like alternate leaf arrange, unilaterally raceme inflorescence and enlarge of adaxial li in the corolla tube (Mayer et.al. 2003).
The ingroup was separated in two sister clades, one consist R. obliquum,
R. spumosum and R. celebicum (Clade A), another consist R. klugioides,
Rhynchoglossum klugioides, R. borneense and R. medusothrix were known have larger size in habit height (character 3), flower length (character 31) and corolla tube length (character 38) than R. obliquum, R. spumosum and R. celebicum. The clade of R. obliquum, R. spumosum and R. celebicum are united by size of the plant from the habit height, size of the leaf, size of the inflorescences, flowers, corollas and the number of stamens. Those three species are known with small flowers, small corollas with narrow lip that tongue-like shape while the other three species (R. klugioides, R. borneense, and R. medusothrix) have larger corollas with lip lower elongate or 3-lobes.
Burtt (1962) make his reconstruction about the affinity among the species of Rhynchoglossum. He suggested that the genus is divided into four groups of morphological characters. First group has large corollas and four equal stamens and all coherent, this is typically for the species known formerly as Klugia (R. azureum and R. notonianum). Second is the group that have large corollas and four fertile stamens with anthers coherent in pairs, one larger and the other smaller. This is typically for R. borneense and R. medusothrix. Third is the group which has large corollas and two fertile stamens coherent with only one species known as R. klugioides. The last group is the species that have small corollas and two fertile stamens typically R. obliquum and R. spumosum.
The result of phylogenetic analysis in this works so far supporting Burtt assumption. On other hand the number of stamens did not make R. klugioides
together in the clade A with R. obliquum and R. spumosum but large corollas have strength character that make the species together with R. borneense and
Malesian species has annual habit and the Klugia clade has perennial habit. (Mayer et.al. 2003).
In the clade A, R. celebicum become as the basal sister to the clade B (R. obliquum and R. spumosum) and supported with a low bootstrap value under 50 %. It is separated with some different characters like leaf shape (character 6) and fruit shape (character 56). Rhynchoglossum obliquum and R. spumosum are allied in one clade. Merrill (1923) placed both species allied because R. spumosum
has smaller habit than R. obliquum and very similar, but both have several same characters like leave shape (character 6), leaf base (character 12), flower length (character 31), calyx tube shape (character 32), stamen number (character 46), and fruit shape (charcter 56) that makes them allied in one clade.
Clade C, R. klugioides becoming as a basal sister to the clade D (R. borneense and R. medusothrix). It is separated with some characters like lamina surface (character 7), calyx tube shape (character 32), number of stamens (character 46) and fruit shape (character 56). In this clade the number of stamens character separate R. klugioides ( 2 stamens) with the other two (4 stamens).
Rynchoglossum borneense and R. medusothrix are allied in one clade (bootstrap value 91 %) with high similarity shown in its several characters like the size of the flower (more than 1.5 cm length), number of stamens (four, coherent in pairs), calyx tube (campanulate) and fruit shape (ovoid). Four number stamens on both species separated those two with other species in Malesia and become an intermediate with another species of Rhynchoglossum which have 4 stamens and formerly identify as genus Klugia (Burtt 1962).
Some phylogenetic analysis shows a low bootstrap value in ingroup clade. It is clear that the morphological analysis needs to be complemented by an analysis of molecular data to provide a better resolution within a clade of
TAXONOMIC TREATMENT
RHYNCHOGLOSSUM Blume
Rhynchoglossum Blume, Bijdr. Fl. Ned.-Ind. 14 (1826) 741 ”Rhinchoglossum”; DC., Prodr. 9 (1845) 275; Benth. In Benth. & Hook. f., Gen. Pl. 2 (1876) 1019; C.B.Clarke, in A.DC & C.DC., Monogr. Phan. 5 (1883) 161; in Hook. f., Fl. Brit. Ind. 4 (1885) 367; Boerl., Handl. Fl. Ned. Ind. 2 (1891) 571; Fritsch in Engl. & Prantl, Natürl. Pflanzenfam. 4, 3B (1895) 156; K. Schum. & Lauterb., Fl. Deutsch. Schutzg. (1901) 542; Koord., Exkursionfl. Java 3 (1912) 195; Merr., Enum. Philipp. Pl. 3 (1923) 454; Schltr., in Engl. Bot. Jahrb. 58 (1923) 298; Ridl., Fl. Mal. Pen. 2 (1923) 539; B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 168; Bakh. f. in Backer & Bakh. f., Fl. Java 2 (1965) 525; A. Weber in Kubitzki & J.W. Kadereit, Fam. Gen. Vasc. Pl. 7 (2004) 128. ― Type:
Rhynchoglossum obliquum Blume.
Antonia R. Br. In Wall., Pl. As. Rar. 3 (1832) 65, auct. non Pohl. ― Type:
Antoniaobliqua (Wall.) R. Br. [= Rhynchoglossum obliquum Blume].
Klugia Schltdl., Linnaea 8 (1833) 248; DC., Prodr. 9 (1845) 275; Benth. In Benth. & Hook. f., Gen. Pl. 2 (1876) 1019; C.B.Clarke, in A.DC & C.DC., Monogr. Phan. 5 (1883) 158; in Hook. f., Fl. Brit. Ind. 4 (1885) 366; Fritsch in Engl. & Prantl, Natürl. Pflanzenfam. 4, 3B (1895) 155. ― Type: Klugia azurea
Schltdl. [= Rhynchoglossum azureum (Schltdl.) B.L. Burtt].
Loxotis, R. Br. ex Benth., Scroph. Ind. (1835) 57; R. Br. In Benn., Pl. Jav. Rar. (1838) 102;.Miq., Fl. Ned. Ind. 2 (1856) 731. ― Type: Loxotis obliqua
(Wall.) R. Br. ex Benth. [=Rhynchoglossum obliquum Blume].
Glossanthus Klein ex Benth., Scroph. Ind. (1835) 57. ― Type:
Glossanthus malabaricus Klein ex Benth. [= Rhynchoglossum azureum (Schltdl.) B.L. Burtt].
infundibuliform, campanulate or urceolate, actinomorphic, adnate at base, sometimes winged; white, pale green to dark green, semi translucence; calyx lobe triangular to acute, 5-merous, entire, segments equal. Corolla tube tubular, zygomorphic, inside glabrous or sparsely yellow dot with puberulent or medussoid pilose hair near mouth; white, bluish to dark purple or azure; tube tubular to cylindric, not swollen, slightly longer than limb ; limb 2-lipped; adaxial lip 2-lobed, shorter and smaller; abaxial lip 3-lobed, seldom undivided, lobes, equal or subequal, apex rounded or elongate to obtuse, larger than adaxial.
Stamens 2 or 4, adnate to corolla tube near middle abaxial limb (if 2 stamens) and adaxial limb (other 2 stamens if 4 stamens); filament flat or terete; anthers basifixed, coherent in pairs, thecae nearly parallel or divaricate, dehiscing longitudinally. Ovary ovoid, 1-loculed, placentas 2, parietal; stigma 1, terminal, subglobose, undivided. Fruit capsule, stalked, ovoid to elongate or capsule-like, smaller (fully closed) or larger (half closed) than calyx remnant, dehiscing loculicidally to base; 2-valvate, straight, not twisted. Seeds minute, cuneate, unappendaged.
Distribution. About 10 species occur in India, Ceylon, South China, Taiwan, Indochina to Malay Archipelago, six species of them found in Malesia; one species in Southern America (Mexico to Peru).
Key to Species of Rhynchoglossum in Malesia
5a. Flower 3-3.5 cm long; calyx lobe 3-5 mm long; larger lip with medussoid pilose hairs on the throat ... R. medusothrix b.Flower 1.8-2.8 cm long; calyx lobe 1-2 mm long; larger lip with
non-medussoid pilose hairs on the throat ...R. borneense
1. Rhynchoglossum borneense Merr. (Fig. 7)
Rhynchoglossum borneense Merr, Univ. Calif. Publ. Bot. 25 (1929) 269; B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 168. ― Type: British North Borneo, Elphinstone Province, Tawao. x.1922-iii.1923. Elmer 21467 (holo PNH n.v.; iso A, BO, K, L, SING).
Herb, fleshy, annuals, non rhizomatous, 40-100 cm height. Stem terete, glabrous, up to 5 mm diameter. Leaves alternate; lamina membranous, ovate to elliptic, oblique, 7-20 cm long by 3.5-10 cm wide; base unequal, one side cuneate to acute, other side rounded to cordate, rise after 2-2.5 cm long from base; apex acuminate with 0.5-1 cm tip; margin entire; nerve distinct; lamina surface glabrous on both surface, pale green; petiole glabrous, terete, 1-2 mm diameter, 2-6 cm long. Inflorescence racemose, spreading, terminal and axillar, opposite with leave, 7-25 cm long, 2 rows up to 10 flowers; main axis glabrous; peduncle glabrous, 1-1.5 cm long; bract absent; pedicel glabrous 0.5-1 cm long; bracteole linier, rise at middle of pedicel, 1-3 mm long. Flower 1.8-2.8 cm long. Calyx tube
campanulate, actinomorphic, white to pale green, semi translucence, adnate at base with line fushion c.4 mm, glabrous, 6-9 mm long and 1-2 mm wide; calyx lobe triangular, entire, glabrous, acuminate 1-2 mm long. Corolla tube tubular, glabrous, smooth, velvety, zygomorphic, bluish to blue lavender, 1-2 cm long, with yellow dot on throat; tube 10-15 mm long; limb 8-10 mm long, 2-lipped; adaxial lip smaller than abaxial, 2 lobes, 3-5 mm; abaxial lip larger than adaxial, tongue-like, obtuse, 5-10 mm. Stamens 4, coherent in pairs; 2 smaller attach at abaxial lip in the middle part; anther ovoid, glabrous, 0.5 mm long; filament flattened, erect, 3 mm long; 2 larger attach at adaxial lip in the middle part; anther oblongish, glabrous, 1 mm long; filament flattened, 2 mm long, twisted. Ovary
3-10 mm long by 3 mm wide; calyx remnant up to 6-8 mm long; stalk glabrous, 0.5-1.4 cm; style remnant 10 -15 mm; Seeds minute, ellipsoid, tessellate 0.3-0.5 mm long.
Distribution. Borneo (Fig. 8)
Habitat. Primary forest and Coral limestone rocks on moist cracks and limestone cliff at 40-300 m.
Notes. Rhynchoglossum borneense is endemic to Borneo. It is allied with
R. medusothrix which both have 4 stamens in flower but different in having of non-medussoid pilose hair in the throat of larger lip on flower tube. Burtt (1962) put this species as an intermediate between the species that formerly as Klugia
(R. ntonianum (Wall.) B.L. Burtt, R. gardneri Theobald & Grupe and R. azureum
(Schltdl.) B.L. Burtt) with the rest of Rhynchoglossum (s.s.) species. When the first time this species described by Merrill (1929), he didn’t sign a number of stamens which is known four that allied with Klugia.
Specimens examined.
Borneo. Tawao, Elphinstone Province. Elmer 21467; Kinabatangan District, Tabin Wildlife Reserve. Poulsen & Banggilon 1672; ibid. Poulsen & Abdilla 1685; Lahad Datu, Tabin. Kiew 5122; East Kutei, G. Tepian Lobang on Menubar River Kostermans 5347; ibid. Kostermans 5414; Sangkulirang, Babi Jolong Aet 649; West Kutai, Kombeng. Endert 5225; Gunong Mendam, North of Tabang
Figure 8 Distribution map of Rhynchoglossum borneense in Malesia in the circle
2. Rhynchoglossum celebicum Karton. sp. nov (Fig. 10)
Rhynchoglosso oblique similis foliis oblongis, fructu capsulari calycis reliquiis parte dimidia inclusa differt. ― Type: Celebes, Resident Menado, OA Poso, Maraowa N. side. 1400-1200 m. 5.viii.1937. Eyma 1572.(holo BO; iso A, K, L).
Fleshy erect herb, annuals, non rhizomatous, about 19-40 cm height. Stem
terete, glabrous 2-5 mm diameter. Leaves alternate, exstipulate; lamina membranous, oblique, oblong 3.5-11 cm long by 1.5-3 cm wide; base unequal, one side rounded, other side cuneate; apex acuminate, tip 0.5 cm long; margin entire; lamina surface glabrous on adaxial side puberulous on abaxial side, pale green; petiole glabrous, terete, 0.5-2 cm long. Inflorescences racemose, spreading, terminal, 9 cm long and lateral 8 cm long, up to 11 flowers; main axis terete, glabrous; peduncle 2-5 cm long; bract absent; pedicel puberulous, 0.5-1 mm long; bracteole linier, glabrous, single 0.5 mm long, rise at the middle of pedicel.
glabrous; anther ovoid, 2-thecae, parallel, opening by longitudinal slit, basifixed, 0.5 mm diameter; filament terete, 2-4 mm long, exerted at the middle of tube on abaxial side. Ovary oblong, glabrous, 1-loculed, 1-3 mm long; style terete, glabrous 1-1.2 cm long; stigma capitate, swollen 1 mm. Fruit capsule-like, elongate, 4-10 mm long by 2-3 mm wide, glabrous; stalk puberulous 2-3 mm long, enclosed by calyx remnant in half; calyx remnant 3-7 mm long; style remnant up to 5-6 mm long; Seeds minute, ellipsoid, tessellate 0.3-0.4 mm long.
Distribution. Sulawesi (Fig. 9)
Habitat. On the open area of mountain forest 1200-1400 m.
Notes. Rhynchoglossum celebicum is the only species in Malesia that have fruit character which is not fully enclosed with calyx remnant. In general sense it is allied with R. obliquum in habit and the shape of flowers but different in the shape of leaves while R. celebicum with oblong leaf shape againt R. obliquum with ovate-elliptic leaf shape. The species is known endemic to Sulawesi island in the middle part.
Specimens examined.
Celebes. Porema Kjellberg 2663; East of Poso District, Maraowa. Eyma 1572.
3. Rhynchoglossum klugioides C.B. Clarke (Fig. 12)
Rhynchoglossum klugioides C.B. Clarke, in A. DC & C. DC., Monogr. Phan. 5 (1883) 163; Merr., Enum. Philipp. Pl. 3 (1923) 454; Elmer, Leafl. Philipp. Bot. 3 (1910) 948; B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 169.
― Type: Luzon, Province of Tayabas. Cuming 824 (holo K; iso BM, K).
Erect fleshy herb, annuals, non rhizomatous, c.35 cm height. Stem terete, glabrous to glabrescent, 3-4 mm diameter; branching nodes not swollen. Leaves
glabrous; style remnant about 10-15 mm long; calyx remnant up to 1 cm long and 0.5 cm wide. Seeds minute, cuneate, ellipsoid, tessellate, 0.4-0.5 mm long.
Distribution. Philippine (Luzon, Catanduanes, Mindoro, Samar, Biliran, Leyte, Mindanao. Fide Merrill (1923)), Moluccas (Ceram). [Fig. 11]
Local Name. Cabeddaya (Bagobo), Pampangod (Philippine)
Habitat. Moist humus covered soil of dense forest and damp forested ravines at 500-2000 m.
Notes.Rhynchoglossum klugioides was previously only known from Philippines islands. In this revision known a new locality in Ceram Island, Moluccas. The species has a large flowers with broader corolla limb than the other species of
Rhynchoglossum and urceolate calyx tube. Also, it has subcoriaceous leaf blade and elongate fruit shape.
Specimen examined.
Moluccas. Ceram. Hatumete-Pas. Kornassi 622; ibid. Eyma 2382. Philippines. Luzon. Camarines Sur. Edano BS76399; Mount Maquiling. Servinas BS16888; Baguio. Elmer 8822; Mt. Alzapan. Ramos & Edano BS45715; Mt. Masingit.
Ramos & Edano BS37560; Irosin (Mt. Bulusan). Elmer 15369; ibid. Elmer 14615; Los Baños (Mt. Maquiling). Elmer 17941; Tayabas. Cuming 824. Catanduanes.
Ramos BS30209. Mindanao. Todaya (Mt. Apo). Elmer 11570.
4. Rhynchoglossum medusothrix B.L. Burtt (Fig. 14)
Rhynchoglossum medusothrix B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 169. ― Type: East Borneo, Berouw, Flatland at base of Mt. Ilas Mapulu, 200 m. 21.ix.1957. Kostermans 13994 (holo L; iso BM n.v., BO, A
n.v., E, L, SING, CANB n.v., P n.v.).
Herb, erect, fleshy, annuals, not rhizomatous, about 40 cm height. Stem
glabrous, terete, 3-4 mm diameter. Leaves alternate; lamina membranous, elliptic, oblique, 6.5-19 cm long by 2.5-10 cm wide; base unequal, one side angustate rise at 0.5-1 cm long from base, other side subcordate; apex acute to acuminate with 0,5 cm tip; margin minutely scabrous; lamina surface glabrous at both side, pale greeen; nerve distinct; petiole glabrous, terete, 0.5-3 cm long. Inflorescence
racemose, spreading, terminal and axillar, opposite with leave up to 20 cm long in terminal and up to 8 cm long axillar, with 10-15 flowers; main axis terete, puberulous; peduncle up to 3 cm long; bract absent; pedicel glabrous, 8-10 mm long; bracteole linier 1-2 mm long, rise from the middle of pedicel. Flower 3-3.5 cm long. Calyx tube campanulate, actinomorphic, 1 cm long and 0.8-1 cm wide, glabrous, white, adnate at base with line fushion 2-2.5 mm long; calyx lobe triangular, acuminate, glabrous, entire, 2-5 mm long. Corolla tube personate, glabrous, zygomorphic, bluish white with blue stripes, inside white, 2-2.5 cm long; tube 2-2.2 cm long, upper part of base with pilose dot; limb 2-lipped, 10-12 mm; adaxial lip smaller than abaxial, 2 lobes, 5 mm long; abaxial lip larger than adaxial, tongue-like, 10-12 mm long by 5 mm wide, throat with medussoid-like pilose hair. Stamens 4, coherent in pairs; 2 smaller, attach at abaxial lip in the middle part; anther ovoid, 1.5 mm long; filament flattened, glabrous, 2 mm long; 2 larger attached at adaxial lip, anther suboblong, 3.5 mm diameter; filament flattened, 2 mm long. Ovary elliptic to ovoid, glabrous, 2-2.5 mm long; style glabrous, terete, 2 cm long; stigma capitate, swollen 1 mm. Fruit ovoid, glabrous, fully closed by calyx remnant, 5-10 mm long by 3-4 mm wide; calyx remnant up to 6-10 mm long; stalk glabrous, 5-15 mm long; style remnant 10-15 mm long.Seeds minute, ellipsoid, tessellate 0.3-0.5 mm long.
Local name. Jambangan Jaja (Tidong Kalabakan)
Habitat. On the limestone area or near moist cave at 200 m.
Notes.Rhynchoglossum medusothrix is only known from Borneo. It is resembles to R. borneense by having 4 stamens in flowers but have a different in having medussoid pilose hair in the throat of larger lip on flower tube. The flowers of
R. medusothrix also larger 3-3.5 cm long than R. borneense 1.8-2.8 cm long.
Specimens examined.
Borneo. Sarawak, Saint Lucia FD. Kadir A2086; Berau, Mt. Ilas Mapulu.
Kostermans 13994.
5. Rhynchoglossum obliquum Blume (Fig. 16)
Rhynchoglossum obliquum Blume, Bijdr. Fl. Ned.-Ind. 14 (1826) 741; C.B.Clarke, in A.DC & C.DC., Monogr. Phan. 5 (1883) 161; in Hook. f., Fl. Brit. Ind. 4 (1884) 367; K. Schum. & Lauterb., Fl. Deutsch. Schutzg. (1901) 542; Elmer, Leafl. Philipp. Bot. 2 (1908) 564; Koord., Exkursionfl. Java 3 (1912) 195; Merr., Enum. Philipp. Pl. 3 (1923) 454; Ridl., Fl. Mal. Pen. 2 (1923) 539; B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 170; Bakh. f. in Backer & Bakh. f., Fl. Java 2 (1965) 525. ― Rhynchoglossum blumei DC., Prodr. 9 (1845) 274 ― Type: Java. Blume 52 (lecto L, sh. no. 909.127-80, designated here).
Wulfenia obliqua Wall., Tent. Fl. Nepal (1826) t.35 ― Loxotis obliqua
(Wall.) Benth., Scroph. Ind. (1835) 57; R. Br., in Benn., Pl. Jav. Rar. (1838) 102, t.24; Miq., Fl. Ned. Ind. 2 (1856) 731, t.35 ―Rhynchoglossum obliquum (Wall.) DC. auct. non Blume, Prodr. 9 (1845) 275 ― Rhynchoglossum obliquum Blume var. parviflorum C.B. Clarke, in A.DC. & C.DC., Monogr. Phan. 5 (1883) 162 ― Type: Nepal. Wallich 407 (holo BM n.v.; iso E).
Loxotis intermedia Benth., Scroph. Ind. (1835) 57 ― Rhynchoglossum obliquum (Wall.) DC. var. intermedium (Benth.) DC., Prodr. 9 (1845) 275 ― Type: Nepal. Wallich 408 (holo BM n.v.).
Rhynchoglossum rheedei DC., Prodr. 9 (1845) 274 ― Type: Rheede Hort. Mal. , 9, t.80
Rhynchoglossum zeylanicum Hook., in Bot. Mag. (1845) t.4198 ― Type: Ceylon. Gardner s.n. (holo BM n.v.).
Rhynchoglossum hologlossum Hayata, Icon. Pl. Formos. 5 (1915) 131 ―
Rhynchoglossum obliquum Blume var. Hologlossum (Hayata) W.T.Wang, Bull. Bot. Res., Harbin 4 (1984) 31 ― Type: Formosa, Rinkiho. viii.1911. Inaba s.n.
(holo TI n.v.).
Rhynchoglossum papuae Schltr., in Engl. Bot. Jahrb. 58 (1923) 299; B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 24 (1962) 171. ― Type: New Guinea, Kaiser Wilhelmsland, bei der Kaulo-Ettappe, 150 m. 8.iv.1904. Schlechter 17524
(lecto BO, designated here).
Erect fleshy herb, annuals, non rhizomatous, about 5-150 cm height. Stem
to 8 cm long, sometimes pendulous, up to 50 flowers; lateral Inflorescences rise at axil leaf of opposite with leaf; main axis terete, puberulous; peduncle 1-4 cm long; bract linier or minute, subtending the inflorescences, puberulous, 2-10 mm long, single; pedicel terete, densely puberulous 1-3 mm long; bracteole linier, glabrous to puberulous, single 3-10 mm long, rise from the middle of pedicel.
Flowers 8-12 mm long. Calyx tube campanulate, actinomorphic,glabrous to glabrescent, pale green to white, 3-5.5 mm long by 1.5-3 mm wide, adnate at base 1.5-4 mm long; calyx lobe triangular, acute or acuminate, 5-merous, 1-2 mm long, white to transparent, margin entire to nearly serrate, puberulous. Corolla tube
tubular, glabrous, zygomorphic, nerved, bilabiate, white, blue to dark purple or whitish blue, yellow pubescent dot at throat, 0.8-1.6 cm long by 0.4 cm wide, nerve reticulate; tube 8-10 mm long; limb 6 mm long, 2-lipped; adaxial lip smaller than abaxial, emarginated or 2 lobes 1-3 mm long; abaxial lip larger than adaxial, undivided, tongue-like or lobate, rounded to nearly trilobe, 3-6 mm long. Stamens
2, coherent in pair, exerted to corolla at the middle of abaxial limb; filament terete or flat 1-5 mm long; anther ovoid to oblong, glabrous, 2 thecae, parallel, opening longitudinally, 0.5-1 mm diameter, basifixed.Ovary ovoid to oblong, superior, glabrous, 1-loculed, 1.5-3 mm long by 1 mm wide; style glabrous 5-10 mm long, terete; stigma capitate, swollen 1 mm diameter. Fruit capsule, ovoid, glabrous, enclosed fully with calyx remnant, shorter than calyx remnant, 2-valvate, connected by the style-base, 3-5 mm long by 2-3 mm wide, dehiscent, opening loculicidally to base; calyx remnant 5-8 mm long; stalk 1-1.5 cm long, glabrous to puberulous; style remnant about 5-8 mm long. Seed minute, cuneate, ellipsoid, tessellate, unappendaged, 0.5 mm long.
Distribution. India, Nepal, South China, Formosa, Indochina, Thailand, Malay archipelago to New Guinea (Fig. 15).
Habitat. Edge of road, open area near small stream, lowland secondary forest, creeping at rocks, Limestone massif with primary forest, remnant forest at 20-1400 m.
Notes. Rhynchoglossum obliquum is the most common species in the genus. It is widespread species ever known. In Malesia is found in every island from Malay Peninsula to New Guinea, including Philippines. It is also found outside Malesian region including India to South China. The species is the type for the genus and having long and crowded inflorescences. It is have a small flower than other species, only 8-12 mm long. It is resembles with R. celebicum but different in leaf blade shape and the fruit. Rhynchoglossum obliquum have ovate to elliptic leaf blade and fully enclosed fruit while R. celebicum have oblong to lanceolate leaf blade and half enclosed fruit. The colour of the corolla tube are varies from purely white to dark blue or purple. Rhynchoglossum papuae from New Guinea placed under synonym because of the similarity in all characters with the type of
R. obliquum and make the distribution of the species till New Guinea.
Specimens examined.
Malay Peninsula. Kelantan, Kuala Betis. Kiew 5259; Kelantan, Dabong. Kiew 5265; Pulau Tioman. Kiew 5207. Sumatra. Mt. Leuser. Sumadijaya 335; ibid. De Wilde & De Wilde-Duyfjes 18043; ibid De Wilde & De Wilde-Duyfjes 19239; ibid. De Wilde & De Wilde-Duyfjes 12254; ibid. De Wilde & De Wilde-Duyfjes 12609; ibid. De Wilde & De Wilde-Duyfjes 19848; ibid. Iwatsuki, Murata, Dransfield & Saerudin S582; Gayoluas. Pringgoatmodjo 107. Deli. Lorzing 14980; Simalungun. Keers 58; Sibolangit. Md. Nur SN7437; ibid. Lorzing 3865; ibid. Lorzing 3865; ibid. Alston 14451; Anai Valley. Kleinhoonte 595; Padang, Panti-Culadak Bünnemeijer 25; Mt. Singgalang. Teijsmann 1175HB; Pajingahan.
Ruttner 67: Muko-muko, Maninjau Lake. Hotta & Okada 1682; Batang Palupuh.
Hotta 25029; Panti Nature Reserve. Van Borssum-Walkees 1762; Antokan river near Pari Panjang. Alston 13763; Sugi Waras. De Voogd 1137; Ranau Lake, Mt. Pakiwang. Van Steenis 3491; Kepahiang. Kasik 97; Mt. Rante Berenong. Iboet 170. Java. Tagogapu. Lorzing 1104; Mt. Burangrang. Backer 14132; Cibeber, Cidadap. Backer 22462; ibid. Bakhuizen 1790; ibid. Bakhuizen 2957; ibid.
Bakhuizen 2158; ibid. Bakhuizen 1760; Mt. Malabar. Backer 26239; Kiara Payung. Backer 23706; Pelabuan Ratu. Koorders 34320ȕ; Jampang Kulon. Van Balgooy 5149; Dago waterfall. Van Steenis 1700; Lembang. Backer12914; Mt. Guntur. Danser 6701; Telaga Remis, Mandiranceng. Vermeulen 12; Ciampea.
